3dCAP-Wheat: An Open-Source Comprehensive Computational Framework Precisely Quantifies Wheat Foliar, Nonfoliar, and Canopy Photosynthesis

Canopy photosynthesis is the sum of photosynthesis of all above-ground photosynthetic tissues. Quantitative roles of nonfoliar tissues in canopy photosynthesis remain elusive due to methodology limitations. Here, we develop the first complete canopy photosynthesis model incorporating all above-ground photosynthetic tissues and validate this model on wheat with state-of-the-art gas exchange measurement facilities. The new model precisely predicts wheat canopy gas exchange rates at different growth stages, weather conditions, and canopy architectural perturbations. Using the model, we systematically study (1) the contribution of both foliar and nonfoliar tissues to wheat canopy photosynthesis and (2) the responses of wheat canopy photosynthesis to plant physiological and architectural changes. We found that (1) at tillering, heading, and milking stages, nonfoliar tissues can contribute ~4, ~32, and ~50% of daily gross canopy photosynthesis (A(cgross); ~2, ~15, and ~-13% of daily net canopy photosynthesis, A(cnet)) and absorb ~6, ~42, and ~60% of total light, respectively; (2) under favorable condition, increasing spike photosynthetic activity, rather than enlarging spike size or awn size, can enhance canopy photosynthesis; (3) covariation in tissue respiratory rate and photosynthetic rate may be a major factor responsible for less than expected increase in daily A(cnet); and (4) in general, erect leaves, lower spike position, shorter plant height, and proper plant densities can benefit daily A(cnet). Overall, the model, together with the facilities for quantifying plant architecture and tissue gas exchange, provides an integrated platform to study canopy photosynthesis and support rational design of photosynthetically efficient wheat crops.